ISSN 0031-0301, Paleontological Journal, 2009, Vol. 43, No. 10, pp. 1316–1329. © Pleiades Publishing, Ltd., 2009. A New Microsporangiate Organ from the Lower Carboniferous of the Novgorod Region, Russia O. A. Orlovaa, N. R. Meyer-Melikian†,a, and N. E. Zavialovab a Moscow State University (MGU), Leninskie gory 1, Moscow, 119991 Russia b Borissiak Paleontological Institute of the Russian Academy of Sciences, ul. Profsoyuznaya 123, Moscow, 117997 Russia e-mail: [email protected] Received February 5, 2008 Abstract—A new species of the genus Telangiopsis, T. nonnae O. Orlova et Zavialova, was described on the basis of a microsporangiate organ from the Lower Carboniferous deposits of the Novgorod Region. The mor- phology of branching fertile axes, synangia, and sporangia was thoroughly studied. The three-dimensional sys- tem of fertile axes branches monopodially; ultimate axes bear numerous connivent bunches of synangia, which consist of three to six basally fused elongated ovate sporangia. The morphology and ultrastructure of prepollen grains were studied, which were extracted from the rock matrix surrounding the sporangia. The two-layered exine includes a well-developed endexine and an alveolate ectexine, with one–three rows of large thin-walled alveolae. The new species was compared with other Early Carboniferous microsporangiate organs. Key words: Early Carboniferous, Novgorod Region, fertile axis, synangia, Lyginopteridales, trilete prepollen, exine ultrastructure. DOI: 10.1134/S003103010910013X INTRODUCTION synangia and numerous casts and imprints of detached synangia were found in yellow ferruginous sandstone. During the three last decades, the interest of bota- In addition to the fertile axes, sterile remains of Lygi- nists dealing with fossil and modern plants to Carbon- nopteridales, Medullosales, and Calamopytiales were iferous synangiate pollen organs has considerably found (Orlova and Snigirevskii, 2003, 2004). Fragmen- increased: earlier studied materials were revised tary fronds of Rhodeopteridium and Sphenopteridium (Millay and Taylor, 1979; Meyer-Berthaud, 1989), and are particularly numerous. The total list of the plant new taxa were described (Eggert and Taylor, 1971; remains found in this locality includes Archaeosig- Stidd et al., 1985; Meyer-Berthaud, 1986; Rowe, 1988; illaria vanuxemii (Goepp.) Kidston, Lepidodendron and others). To date, more than 20 genera of microspo- acuminatum L. veltheimii Lepi- rangiate organs have been described from the Carbon- (Goepp.) Stur, Sternb., dostrobus Lepidostrobus ornatus Flem- iferous deposits of Europe, Great Britain, and North sp. 1, Brongn., and South Americas (Erwin et al., 1994). They belong ingites sp. 1, Stigmaria sp., Archaeocalamites radiatus to four orders of Lyginopteridophyta: Calamopytiales, (Brongn.) Stur, Sphenopteridium bifidum (L. et H.) Lyginopteridales, Callistophyllales, and Medullosales. Benson, Sphenopteridium sp. 1, S. pachyrrhachis Early Carboniferous deposits only contain members of (Goepp.) Pot., S. jurinae O. Orl. et S. Snig., S. gaebleri the first and second orders. The majority of them were Gothan, Adiantites antiquus (Ett.) Stur, A. machanekii found in Scottish and English localities, and the rest Stur, Adiantites sp. 1, Sphenopteris distans Sternb., was reported from the United States, Germany, and Sphenopteris sp. 1, S. foliolata Stur., Rhodeopteridium Russia (Orlova, 2002, 2007). The first Russian find of a hochstetteri (Stur) Purk., R. goeppertii (Ett.) O. Orl. et synangiate organ was preliminary discussed by Orlova S. Snig., R. tenue (Gothan) Kotas., Lyginodendron sp., (2002); the present paper publishes a detailed study of and Rhynchogonium sulcatum (L. et H.) Zal. (Orlova this fossil. and Rasskazova, 2005). The deposits of the Porog Vittsa 2 locality are char- acterized by a palynological assemblage with con- MATERIAL AND METHODS stantly occurring numerous Lycospora pusilla (Ibr.) The material was collected from the Porog Vittsa 2 Som. Lophotriletes grossepunctatus (Waltz) Isch., locality, situated at the right bank of the Msta River, Triquitrites marginatus H., St. et M., T. comptus Will., near the village of Putlino, Borovichi district of the Trachytriletes subintortus Isch., Schulzospora campy- Novgorod Region. An imprint of a fertile axis bearing loptera (Waltz) H., St. et M., S. magnifica (Isch.) 1316 A NEW MICROSPORANGIATE ORGAN FROM THE LOWER CARBONIFEROUS 1317 Fig. 1. Telangiopsis nonnae sp. nov., holotype MGU, no. 293/47; general view of microsporangiate organ, branching fertile axes and clusters of synangia are shown, ×3.5. Porog Vittsa 2 locality, Borovichi district, Novgorod Region, Russia; Lower Carbonifer- ous, Visean Stage, Upper Visean Substage. Oshurk., Tripartites vetustus Schem., Punctatisporites Collection, no. 293 is kept at the Department of glaber (Naum.) Playf., and some other spores occurring Paleontology, MGU. Olympus SZ-6045 and Leica much more rarely (Orlova and Rasskazova, 2005). The MZ16 stereomicroscopes were used during the study palynological assemblage fits the palynological zones and photographing. A rock fragment with detached Perotriletes tessellatus–Schulzospora campyloptera synangia was macerated after Grishchuk (fluoric acid (TC), Raistrickia nigra–Triquitrites marginatus (NM), followed by centrifugation). The organic residue with Tripartites vetustus–Rotaspora fracta (VF), and Bel- prepollen grains was mounted on slides and studied in lispores nitidus–Reticulatisporites carnosus (NC) of the transmitted light, under a Zeiss Axioplan 2 microscope; Upper Visean=Namurian A (Serpukhovian) of West some prepollen grains were studied under Camscan and Europe. Hitachi S-405A scanning electron microscopes PALEONTOLOGICAL JOURNAL Vol. 43 No. 10 2009 1318 ORLOVA et al. Carboniferous, Visean Stage, Upper Visean Substage (Fig. 1). Diagnosis. System of fertile axes branches monopodially, three-dimensional. Main axis with alter- nating secondary axes. Secondary axes bear alternating tertiary axes terminating with ultimate axes of numer- ous connivent clusters of synangia gathering in pairs, or, occasionally, single. Synangia 1.2–3.5 mm long and up to 3.5 mm wide in terminal part. Each synangium consists of three–six elongated ovate basally fused spo- rangia. External surface of sporangia with coarse longi- tudinal ribs. Prepollen circular to circular-triangular and oval, 27.8 x 32.0 μm, trilete, rays reach 3/4–4/5 of radius. Exine surface verrucate, folded, and equatori- ally echinate. Exine bilayered, considerably varying in thickness, with alveolate ectexine. Description (Figs. 1–4). The leafless fertile axis shows a compound architecture. Fertile axes of all orders alternate in three dimensions. The visible length of the main axis (=rachis) is 34 mm, and the width is 2.5 mm. The rachis bears alternating lateral branches (secondary axes), which are situated at a distance of 4–11 mm from each other. The upper axis of the second order is only preserved in the studied specimen (Pl. 25, fig. 2; Pl. 26, fig. 1; Figs. 1, 2). The next secondary axis was situated at a distance of 11 mm below the visible secondary axis. The preserved secondary axis deviates from the main axis at an angle of about 35° and branches three- Fig. 2. Reconstruction of pollen organ of Telangiopsis non- nae sp. nov. Scale bar 2 mm. dimensionally, producing alternating branches. The width of this secondary axis is 0.5–0.6 mm. The tertiary axis deviates from the secondary axis at an angle of 45° (SEMs). Ultrathin sections were made using an LKB 3 at a distance of 5 mm from the point where the second- ultramicrotome with a diamond knife and studied under ary axis deviates from the main axis. The terminal fer- a Hitachi H-600 transmission electron microscope tile axis of the forth order bearing synangia deviates (TEM). In total, 21 prepollen grains were studied in from the tertiary axis (Pl. 25, fig. 2). All axes are cov- transmitted light; 18, under SEM; and 5, under TEM. ered with longitudinal ribs (Pl. 26, fig. 1). The distance Grids with ultrathin sections and films and graphical between adjacent ribs is 0.15–0.2 mm. files documenting the sections are kept at the laboratory The synangia deviate from the ultimate axis in three of paleobotany, PIN. dimensions. The synangia are attached to the axis with short petioles. Most often, the synangia are in pairs; or, SYSTEMATIC PALEOBOTANY more rarely, they are solitary (Fig. 2). Usually, the clus- ters of synangia are situated closely to each other and Division Lyginopteridophyta often are partially or completely superimposed (Pl. 25, Class Lyginopteridopsida fig. 1). The synangia consist of three to six sporangia Subclass Lyginopterididae (Fig. 3). Most often, synangia of four or five sporangia Order Lyginopteridales occur (Figs. 3c–3e; Pl. 26, figs. 2, 4). Usually, synangia Family Lyginopteridaceae Potonie, 1902 are situated along the bedding plane, and, therefore, in longitudinal section resembling wide cups (Figs. 2, 3; Genus Telangiopsis Eggert et Taylor, 1971 Pl. 26, figs. 2–6). Rarely, they are tear-shaped, small, Telangiopsis nonnae O. Orlova et Zavialova, sp. nov. and consisting of three sporangia (Fig. 3b). Most prob- Plates 25–29 ably, such synangia were immature. The length of the Etymology. In memory to Nonna Robertovna sporangia preserved along the bedding plane varies Meyer-Melikian, our scientific teacher and outstanding from 1.2 to 3.5 mm; and the width in the terminal part botanist, died an early death. reaches 3.5 mm (Table 1). Some synangia are pros- Holotype. MGU, no. 293/47, Geological